Identification of Single Nucleotide Polymorphism of Adipocyte
Fatty Acid-Binding Protein Gene and Its Association
with Fatness Traits in the Chicken1
Q. Wang,* H. Li,*2 N. Li,† L. Leng,* Y. Wang,* and Z. Tang*
*College of Animal Science and Technology, Northeast Agricultural University, Harbin 150030, People’s Republic of China;
and †National Laboratories for Agribiotechnology, China Agricultural University, Beijing 100094, People’s Republic of China
ABSTRACT Fatty acid-binding proteins (FABP) belong
to a superfamily of lipid binding proteins that exhibit a
high affinity for long chain fatty acids and appear to
function in metabolism and intracellular transportation
of lipids. The current study was designed to investigate
the effects of the adipocyte (A)-FABP gene on chicken
growth and body composition traits. Two F2 resource
populations were used in the current study. Body weight
and body composition traits were measured in the popu-
lations. Primers for the coding region of the A-FABP gene
Key words: A-FABP gene, growth, body composition, polymorphism, chicken
INTRODUCTION
In the past half century, selection in broiler chickens
for increased growth rate and, as a consequence, reduced
market age have greatly increased feed efficiency; how-
ever, the modern broilers selected for rapid growth ex-
hibit higher body fat deposition (Mallard and Douaire,
1988; Griffin, 1996). Fat is considered a by-product with
very low commercial value. It is not a cost-effective body
component from a view of energy use, and large amounts
of fat deposition can depress feed efficiency. Although
several strategies of selection for leanness in poultry pro-
duction have been described, it is still difficult to measure
body fat content (Mallard and Douaire, 1988; Jennen et
al., 2004). The measurements of fatness traits (for example,
abdominal and skin fat content) are tedious and expen-
sive. Molecular genetic information is required to be used
to enhance genetic improvement of animal species. Identi-
fying the QTL responsible for the economic important
traits will facilitate poultry breeding programs. There are
2 basic methods of QTL identification, the candidate gene
approach and the whole-genome scanning (Rothschild
2006 Poultry Science Association, Inc.
Received July 22, 2005.
Accepted November 17, 2005.
1
This research was supported by National 863 Project (No.
2002AA211021) and National Natural Science Funds (No. 30270950).
2
Corresponding author: lihui@neau.edu.cn or lihui645@hotmail.com
429
were designed from chicken genomic and cDNA se-
quences. Polymorphism between parental lines was de-
tected by DNA sequencing, and the PCR-RFLP method
was developed to genotype the F2 populations. The re-
sults indicated that an A-FABP gene polymorphism was
associated with abdominal fat weight and percentage of
abdominal fat, and the A-FABP gene could be a candidate
locus or linked to a major gene(s) that affects abdominal
fat content in the chicken.
2006 Poultry Science 85:429–434
and Soller, 1997; Dodgson and Cheng, 1999). The candi-
date gene approach is an economic method to find QTL
responsible for genetic variation in traits of interest (Roth-
schild and Soller, 1997).
Fatty acid-binding protein (FABP) genes are important
genes that are involved in the development of fatness
traits (McArthur et al., 1999) and that belong to a super-
family of lipid-binding proteins and occur intracellularly
in invertebrates and vertebrates. The FABP are small mo-
lecular weight proteins with high binding affinities for
long chain fatty acids (FA; Ockner et al., 1972). These
proteins are involved in intracellular fatty acid transpor-
tation (Tipping and Ketterer, 1981), cell growth and differ-
entiation, cellular signaling, gene transcription, and
protection of enzymes from the toxic effects of free FA
(Glatz and Veerkamp, 1985; Zimmerman and Veerkamp,
2002). They may also play a role in the modulation of
enzyme activity and signal transduction (Grinstead et
al., 1983).
In mammals, the intracellular or cytoplasmic FABP
form a group of at least 9 distinct proteins, which are
liver FABP, intestinal FABP, heart FABP, adipocyte
(A−) FABP, epidermal (E−)FABP, ileal FABP, brain FABP,
myelin FABP, and testicular FABP (Zimmerman and
Veerkamp, 2002). Their molecular weight ranges from 14
to 15 kDa with 126 to 134 amino acids and are named
for the tissues from which they are isolated (Ockner et
al., 1972). Some tissues express several types, either in
different cell types (brain, kidney, and stomach) or in
430
Figure 1. The PCR-RFLP pattern for A-FABP gene with TaqI digestion. A = granddam (Baier layer and Silky) allele; B = grandsire (broiler)
allele; M = marker DL2000. The fragment sizes of the marker are listed on the left side.
the same cells (enterocyte). All FABP types show similar
structural features containing 10 antiparallel β strands
(βA-βJ) that form a β barrel. The bound ligand is found
within the barrel in a central internal water-filled cavity.
The overall organization of all members of the FABP
family is identical, containing 4 exons and 3 introns. The
exon/intron positions are similar, but the intron length
is variable among the genes (Zimmerman and Veer-
kamp, 2002).
The A-FABP gene has been cloned and mapped in
humans, mice, pigs, and chickens (Hunt et al., 1986;
Prinsen et al., 1997; Gerbens et al., 1998; Wang et al., 2004).
This gene is expressed only in the fat tissue (Fisher et al.,
2001; Wang et al., 2004). Mice lacking A-FABP are healthy,
develop apparently normal adipose tissue, and exhibit
only minor alterations in their steady-state lipid metabo-
lism (Hotamisligil et al., 1996). This lack of significant
effect of A-FABP deficiency was potentially the conse-
quence of the compensatory upregulation of E-FABP
mRNA (Coe et al., 1999; Shaughnessy et al., 2000). There
was no difference in the rate of FA influx or esterification
in adipocytes of wild-type and A-FABP null mice, but
basal lipolysis was decreased approximately 40% in A-
FABP null mice (Coe et al., 1999; Jenkins-Kruchten et al.,
2003). Porcine A-FABP gene variation within the first
intron is associated with differences in intramuscular fat
content and possibly growth in a Duroc population (Ger-
bens et al., 1998). It was reported recently that the poly-
morphism, A-FABP4-A376C and PPAR gamma
Pro12Ala, worked together to influence a biologic path-
way affecting insulin sensitivity and body composition
in nondiabetic Hispanic and nonHispanic white males
(Damcott et al., 2004). The objectives of the present study
were to identify polymorphism of the chicken A-FABP
gene in the resource populations, to develop a PCR-RFLP
method for the genotyping, and to evaluate the associa-
tions of the polymorphism of the gene with growth and
body composition traits.
WANG ET AL.
MATERIALS AND METHODS
Resource Populations
Two populations, Northeast Agricultural University
Resource Population (NEAURP) and China Agricultural
University Resource Population (CAURP), were used in
the current study. The NEAURP was established by cross-
ing broiler sires, derived from lines at NEAU divergently
selected for abdominal fat, with Baier layer dams, a Chi-
nese local breed. The F1 birds were intercrossed to pro-
duce F2 offspring. A total of 455 F2 individuals were
produced from 5 F1 families for the study. The CAURP
was established by crossing broilers with Silkies, and 558
F2 individuals were produced from 4 families for the
study. All birds had free access to feed and water. Com-
mercial corn-soybean-based diets that met all NRC re-
quirements (National Research Council, 1994) were
provided in the study. From hatch to 3 wk of age, birds
received a starter feed (3,000 kcal of ME/kg and 210 g/
kg of CP) and from 3 to 12 wk of age, birds were fed a
grower diet (3,100 kcal of ME/kg and 190 g/kg of CP).
Phenotyping
Live BW was measured at hatch and weekly up to 12
wk of age. Chicken body composition traits were re-
corded after slaughter at 12 wk of age. These measure-
ments included pectoralis major weight (PMaW),
pectoralis minor weight (PMiW), leg muscle weight
(LMW), abdominal fat weight (AFW), spleen weight
(SW), liver weight (LW), and heart weight (HW). All the
traits were also expressed as a percentage of BW at 12
wk of age.
DNA and RNA Extraction
and cDNA Synthesis
Genomic DNA was isolated from venous blood sam-
ples. Twenty microliters of blood samples were collected
 CHICKEN A-FABP GENE AND FATNESS TRAITS 431
Table 1. Effects (P-value) of adipocyte fatty acid-binding protein poly- centrifugation at 10,000 rpm for 10 min. The DNA was
morphisms on chicken growth and body composition traits
washed with 70% ethanol and resuspended in 200 ␮L of
Population2 TE buffer (10 mM Tris-HCl and 1 mM EDTA, pH 8.0).
Trait1 Age (wk) NEAURP CAURP Total RNA was isolated from 3 grandsires and 3 grand-
3
dams (2 from each genetic cross). Adipose tissue samples
BW 0 NS NS
BW 1 NS 0.1369
were frozen in liquid nitrogen and pulverized with a
BW 2 NS 0.0108 mortar and a pestle. The RNA was extracted with the
BW 3 NS 0.0105 Trizol reagent (Invitrogen, Rockville, MD) according to
BW 4 0.0736 NS
BW 5 0.1656 0.0134
the instructions of the manufacturer, then resuspended
BW 6 0.1114 0.0261 in diethyl pyrocarbonate-treated water. The reverse tran-
BW 7 0.1349 0.0784 scription reactions were carried out using the TaKaRa
BW 8 NS 0.0117
BW 9 NS 0.0094
RNA PCR Kit Ver.2.1 (TaKaRa Biotechnology Co., Ltd.,
BW 10 NS 0.0170 Dalian, China).
BW 11 NS 0.0014
BW 12 NS 0.0001
CW 12 NS 0.0091 DNA Sequencing of Chicken A-FABP Gene
PMaW 12 NS 0.0515
PMiW 12 0.0392 NS Primers for amplifying a fragment of 434 bp from the
LMW 12 NS 0.0246
LW 12 NS 0.0870
A-FABP cDNA were AFF, 5′-AGA CTG CTA CCT GGC
AFW 12 0.0048 0.0224 CTG ACA-3′; and AFR, 5′-AAG ACG GCT TCC TCA
%PMaW 12 NS NS TGC-3′. The PCR components were 1× PCR reaction
%PMiW 12 0.0019 NS
%LMW 12 NS NS buffer (10 mM Tris-HCl, 50 mM KCl, and 1.5 mM MgCl2,
%LW 12 NS NS pH 8.3), 200 ␮M of each deoxynucleotide triphosphate,
%AFW 12 0.0014 0.0778 0.25 ␮M of each primer, 2 ␮L of cDNA, and 1 U of Taq
1
CW= carcass weight; PMaW = pectoralis major weight; PMiW = DNA polymerase (TaKaRa Biotechnology Co.) in a final
pectoralis minor weight; LMW= leg muscle weight; AFW = abdominal volume of 25 ␮L. The thermal cycling were 94°C for 5
fat weight; LW = liver weight; and % indicates that traits are expressed
as percentage of BW at 12 wk of age.
min followed by 30 cycles of 94°C for 50 s, 55°C for 1
2
NEAURP = Northeast Agricultural University Resource Population; min, and 72°C for 1 min. The PCR products were purified
CAURP = China Agricultural University Resource Population. by Gel Extraction Mini Kit (Watson Biotechnologies, Inc.,
3
NS = P > 0.2. Shanghai, China) and sequenced in forward and reverse
directions with an ABI 3730 sequencer (Bioasia Biotech-
nology Co., Ltd., Shanghai, China). Restriction enzyme
in EDTA-Na2-coated tubes, mixed with 445 ␮L of SET sites in these sequences were detected by the DNAMAN
buffer (50 mM Tris, 150 mM NaCl, and 1 mM EDTA), 25 package Version 4.0 (Lynnon BioSoft, Vaudreuil-Dorion,
␮L of SDS (10%), and 10 ␮L of proteinase K (10 mg/mL). Quebec, Canada).
This mixture was incubated at 56°C for 8 h, and then 200
␮L of NaCl (5 M) was added to the lysate. The solutions Genotyping of Chicken A-FABP Gene
were mixed by vortex for 10 min. Subsequently, the mix-
ture was centrifuged at 12,000 rpm for 10 min, and the A genomic fragment of 702 bp including exons 1 and
supernatant was transferred to a new tube and added partial intron 1 was amplified by using primers AFF (5′-
with an equal volume of chloroform. Then, the mixture AGA CTG CTA CCT GGC CTG ACA-3′) and AFR2 (5′-
was centrifuged again at 12,000 rpm for 10 min after CAT CCT ACT GGA ATA CGG-3′). The PCR components
mixing by vortex for 10 min. Genomic DNA was precipi- included 1× PCR reaction buffer, 200 ␮M of each deoxy-
tated with twice volume of 100% ethanol followed by nucleotide triphosphate, 0.25 ␮M of each primer, 1 ␮L

Table 2. Effects (least square means) of adipocyte fatty acid-binding protein genotype on body composition
traits at the Northeast Agricultural University Resource Population

Genotype2 (n)
Trait1 AA AB BB

BW4 (g) 435.06 ± b

5.51 (113) 443.09 ± ab
4.00 (220) 453.16 ± 5.70a (107)
AFW (g) 73.90 ± 2.57b (115) 84.08 ± 1.85a (227) 82.57 ± 2.62a (113)
%AFW (g/100 g of BW) 3.72 ± 0.106b (115) 4.19 ± 0.077a (227) 4.09 ± 0.109a (113)
PMiW (g) 54.71 ± 1.01b (114) 55.66 ± 0.72b (227) 58.19 ± 1.02a (112)
%PMiW (g/100 g of BW) 2.70 ± 0.028B (114) 2.75 ± 0.020B (227) 2.84 ± 0.029A (112)
Means within a row with no common superscript are different (P < 0.05).
a,b

Means within a row with no common superscript are different (P < 0.01).
A,B
1
BW4 = BW at 4 wk of age; AFW = abdominal fat weight; PMiW = pectoralis minor weight; and % indicates
that traits are expressed as a percentage of BW at 12 wk of age.
2
Numbers showed in parentheses are the individuals selected.
432 WANG ET AL.
Table 3. Effects (least square means) of adipocyte fatty acid-binding protein genotype on body composition
traits at the China Agricultural University Resource Population

Genotype2 (n)

Trait (unit)1 AA AB BB
BW1(g) 61.65 ± b
3.99 (32) 65.95 ± a
3.53 (185) 65.75 ± 3.51a (302)
BW2(g) 125.72 ± 7.27b (29) 137.93 ± 6.20a (182) 134.39 ± 6.14ab (300)
BW3(g) 213.79 ± 12.26b (31) 232.28 ± 10.62a (184) 225.30 ± 10.53b (309)
BW5(g) 452.87 ± 26.34B (31) 502.46 ± 21.97A (181) 499.31 ± 21.74A (284)
BW6(g) 627.37 ± 25.05b (28) 693.58 ± 12.33a (172) 679.86 ± 11.41a (274)
BW7(g) 809.35 ± 32.01b (28) 880.12 ± 15.12a (186) 865.15 ± 13.88ab (297)
BW8(g) 963.11 ± 39.56B (30) 1,076.69 ± 22.01A (172) 1,071.59 ± 20.68A (281)
BW9(g) 1,155.30 ± 41.87B (32) 1,281.18 ± 22.57A (175) 1,274.84 ± 20.76A (295)
BW10(g) 1,321.61 ± 52.59B (28) 1,458.51 ± 31.98A (172) 1,453.86 ± 30.85A (273)
BW11(g) 1,408.61 ± 67.04B (23) 1,618.14 ± 44.20A (152) 1,581.48 ± 42.73A (247)
BW12(g) 1,523.45 ± 62.28B (26) 1,759.59 ± 38.92A (167) 1,713.36 ± 37.37A (273)
CW(g) 1,373.31 ± 53.71b (35) 1,516.29 ± 35.70a (200) 1,496.41 ± 34.53a (324)
PMaW (g) 85.99 ± 4.73b (35) 95.46 ± 3.35a (200) 94.48 ± 3.27a (324)
LMW(g) 119.73 ± 5.54b (35) 133.74 ± 3.34a (200) 132.38 ± 3.19a (324)
LW(g) 31.03 ± 4.09b (35) 34.06 ± 3.91a (200) 34.02 ± 3.90a (323)
AFW (g) 35.46 ± 6.69b (35) 49.21 ± 5.12a (200) 48.47 ± 5.04a (323)
%AFW (g/100 g of BW) 2.60 ± 0.38b (35) 3.22 ± 0.30a (200) 3.26 ± 0.28a (323)
a,b
Means within a row with no common superscript are different (P < 0.05).
A,B
Means within a row with no common superscript are different (P < 0.01).
1
BW1, BW2, BW3, BW5, BW6, BW7, BW8, BW9, BW10, BW11, and BW12 = BW at 1, 2, 3, 5, 6, 7, 8, 9,
10, 11, and 12 wk, respectively; CW = carcass weight; PMaW = pectoralis major weight; PMiW = pectoralis
minor weight; LMW = leg muscle weight; LW = liver weight; AFW = abdominal fat weight; and % indicates
that traits are expressed as a percentage of BW at 12 wk of age.
2
Numbers shown in parentheses are the individuals selected.

of genomic DNA (50 ng/␮L), and 0.5 U of Taq DNA
polymerase in a final volume of 10 ␮L. The thermal pro-
files were 94°C for 5 min followed by 32 cycles of 94°C
for 30 s, 58°C for 30 s, and 72°C for 50 s. A single nucleotide
polymorphism (SNP) of the A-FABP gene was detected
by digesting 7 ␮L of the 702-bp PCR product with 5 U
of the TaqI enzyme (TaKaRa Biotechnology Co.) at 65°C
overnight. Restriction patterns were visualized by agar-
ose gel electrophresis and ethidium bromide staining.
Statistical Analysis
Data were subjected to the GLM procedures of JMP
(SAS Inst. Inc., Cary, NC) with genotype (G) and sex (S)
as fixed effects, family (f) and hatch (h) as random effects,
according to the model: Y = ␮ + G + S + f + h + e; here
Y was the dependent variable, ␮ was population mean,
and e was the random error. The interaction G × S was
not significant for any trait and therefore was not included
in the model. Statistical significance threshold was deter-
mined as P < 0.05, unless otherwise specified.
RESULTS
Identification of Polymorphism
and PCR-RFLP Analysis
The aligned complete coding sequences of the chicken
A-FABP gene revealed 1 SNP (a C/T substitution), which
was located in the exon 1 of the A-FABP gene. It was
a silent mutation compared with a GenBank sequence
(AF432507). This mutation was associated with a TaqI
restriction site. The 702-bp fragment was digested with
TaqI restriction enzyme. The broiler allele that was desig-
nated as B displayed a band pattern of 2 fragments (629
and 73 bp), and the layer allele or A allele showed only
1 fragment (702 bp; Figure 1).
Association of A-FABP Gene SNP
with Growth and Body Composition
The genotypes of the SNP of the chicken A-FABP gene
were used in the genetics analysis of the resource popula-
tion. There was a significant association (P < 0.05) between
the SNP and body composition traits (PMiW, %PMiW,
AFW, and %AFW) in F2 birds of the NEAURP (Table 1).
There were significant associations (P < 0.05) between the
SNP and BW at 2, 3, 5, 6, 8, 9, 10, 11, and 12 wk of age
and body composition traits (CW, LMW, and AFW) in
F2 birds in the CAURP.
Effects (least square means) of the A-FABP genotype
on growth and body composition traits in F2 offspring of
the NEAURP and the CAURP are shown in Tables 2
and 3, respectively. At the NEAURP location, there were
lower (P < 0.05) BW at 4 wk of age in F2 birds that were
of the AA genotype than those of the BB genotype; there
were lower (P < 0.01) AFW and %AFW in F2 birds that
were AA genotype than those of AB and BB genotypes;
there were lower PMiW (P < 0.05) and % PMiW (P < 0.01)
in F2 birds that were AA and AB genotypes than those
of the BB genotype. At the CAURP location, there were
lower (P < 0.05) BW at 1, 5, 6, 8, 9, 10, 11, and 12 wk of
age and CW, PMaW, LMW, LW, AFW, and %AFW in F2
birds that were of the AA genotype than those of the AB
and BB genotypes; there were lower (P < 0.05) BW at 2
and 7 wk of age in F2 birds that were of the AA genotype
 CHICKEN A-FABP GENE AND FATNESS TRAITS
than those of the AB genotype; there were lower (P <
0.05) BW at 3 wk of age in F2 birds that were of the AA
and BB genotypes than those of the AB genotype.
DISCUSSION
The FABP specific occurrence in certain tissues or cells
possibly results from adaptation to specific cellular needs
(Zimmerman and Veerkamp, 2002). In the present study,
the detected mutation within the chicken A-FABP gene
was located in exon 1. Statistic analysis indicated that the
SNP was related with growth (BW) and body composition
(AFW and AFW%) traits.
Growth is a comprehensive reflection of development
of various parts of the chicken body, and its final expres-
sion is the result of interaction among genetic, nutritional,
and environmental factors (Scanes et al., 1984). Growth
is under complex genetic control, and uncovering the
molecular mechanisms of growth will contribute to more
efficient selection for growth in broiler chickens. In the
present study, the A-FABP SNP had an association (P <
0.05) with BW in the CAURP. The F2 birds with the A-
FABP-AA genotype had a lower BW than the birds with
the A-FABP-BB and -AB genotypes. In an F2 cross between
outbred lines, however, the linkage disequilibrium is sub-
stantial. The associations observed among A-FABP-SNP
and BW at different weeks of age in the CAURP might
have been produced by linkage disequilibrium between
the SNP and another mutation located in the A-FABP
locus or another linked gene directly involved in the regu-
lation of these phenotypic traits. This interpretation is
more consistent with the current data because it may
explain why no associations were found between A-
FABP-SNP genotypes and BW in both F2 populations
simultaneously. The SNP can be interpreted as a molecu-
lar marker linked with a major gene to significantly affect
growth traits in chicken.
Excessive fat deposition in chickens should be limited
to enhance production efficiency and product quality.
Similar to the human, pig, and mouse A-FABP genes, the
chicken A-FABP gene is expressed only in fat tissue, in
which it is extremely abundant. The A-FABP gene is,
therefore, a logical target for investigation of its effects
on chicken fat metabolism (Wang et al., 2004). The current
study found that a polymorphism of exon 1 region of the
A-FABP gene was related (P < 0.05) with AFW and %AFW
at 12 wk of age in both F2 populations. The F2 birds with
the A-FABP-AA genotype had a lower AFW and %AFW
than the birds with the A-FABP-BB genotypes. Compari-
son of mean values for all 3 genotypes suggests that A-
FABP acts in an additive fashion on fatness traits with
the broiler allele contributing to greater AFW and %AFW.
The allele association (broiler with more fat content in
the body) found in the present study was consistent with
the selection history of broilers, emphasizing growth rate
to market age that lead to more fat deposition in the body.
In an F2 cross of divergent lines, the linkage disequilib-
rium is substantial, and the examined SNP may be causal
or may be linked with functional polymorphisms of A-
433
FABP or influence the expression of the gene or be linked
to a major gene and therefore affect expression of abdomi-
nal fat content.
One of the breeding goals for broilers is to decrease
abdominal fat content. The measurement of the trait is
laborious and expensive by slaughtering birds based on
traditional selection methods. Therefore, molecular MAS
can improve genetic selection programs. The results from
the current study indicated that a SNP marker in the A-
FABP gene was associated with abdominal fat content in
chickens and is, therefore, a potential marker for use in
molecular MAS programs.
ACKNOWLEDGMENTS
The authors gratefully acknowledge the members of
the Poultry Farm located at Northeast Agricultural Uni-
versity for managing the birds. The authors are also in-
debted particularly X. Deng and coworkers.
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